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305-8572 ȏòǒ#-ġĂƸÓ 1ɼ1ɼ1Biogeography of a water-bloom forming cyanobacterium Microcystis aeruginosa
Key words: biogeography; FST Microcystis; MLST; waterbloom.
Yuuhiko Tanabe
Graduate School of Life & Environmental Sciences, University of Tsukuba
1-1-1 Tennodai, Tsukuba, Ibaraki 305-8572, Japan/.  '$&3 ƽƳïƻĉ,ïƺ* >ƽƳǞ+ǡɕ¶ĢBȫũ>7+ĉã& > (Avise 2000)'@&ɻ~ȍ*ɶǦÁƳ9āñƂƳBĔȱ'% >ǗǠȀ+ƽƳïƻĉ+ǗǠŦƝ,ɻƇ)ïó*% >Ǟ > ,+ȼǻǞ+¶ĢBȬ1ɻĀŠ+¤BŔɢ%ĲņȟēǦBȔ ɻɈı)ɊĆo}K}B"%ǮǸ|ɢçȠŻBȔ"Ƿż'ïƻ¶Ģ'+ɖ¡BȰȭ>ɻ'8+&,)@ gO`vE*# %823×Ƈ+Ely}]BȷȚ>'*;"%ƽƳïƻĉǗǠȔA?%%, ɻƑľ)<+Ƿż,ɶǦÁƂƳ+ǗǠŇż*Ȝ<?>; )OvE)8+&,)ɻ)=+ŰŭB 8+& "+Éæ,gO`vE
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Everythingis everywhere: where environment selectsɹńȥʄĺƽƳ+ǡɕ¶Ģ,Ƽúțæ*;"%+5ƙČ?>ɺ'ØÑ&ȗƹ (Baas Becking 1934)+ȫ,Ƿę+'@ɻgO`vE+ƽƳïƻĉ'ǗǠ¶ɐ,88ŇǣĹ) ɹǗǠ'@&ïƻǌh[}z,µ%)<ńÞ)

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Y. Tanabe - 1

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Y. Tanabe - 2

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Y. Tanabe - 3ŷǞ, 3-8m ǝĪ+TGY+ÇǲȄŀ+ƺȐ& >ɻȿĥ,¤+ǲȄɢÖɻĒĂȳBÜC Ĳņǌ*ĀƇ)Syd}BĲŇ>'&Ǖ<?>Ð,+ĀƇ)Syd}Ĳņ*;"%ʁ#+Ǟ*¶ɰ?% ɻɊĆxmw&2'C(ɇ ) 'ɻSyd}Ĳ
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Y. Tanabe - 4

MLSTɹMultilocus sequence typingɺ'(Maiden et al. 1998)ŷŉƝƗƿ?% >ƻǁ'%ɻgO`vE,~ȍ*ɶǦÁƂƳǦ'Ɣȹ%Ǟ±+ɊǌĀƇŀāɻ[zhOȳBS
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Y. Tanabe - 5

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Ň+ǘƷȭǌýÁɹ§ư*ĆĊBƚĝƑ=ɻƑ)"='ƵƜBƽ6Àżɺ*;"%ɻ#+ɢçɕ&ɊĆñɬĪɹƴČ+ɊĆñ+ɢç*È7>ɬĪɺ,ýA"% +ƒɜɹéɧɺ&,FST 'ɊĆñɬĪBýŠ'>ŒŠBƿ >'*;"%ɻɢçɕ+ɊǌǵŇ+ɇ +ŴƯ+ƃµɹ0 ;=Ŵń*ā?-ǵŇ,ǆ)>'¹ŤɺɻË.ɊǌǵŇ+ā+ɇ BŖƩɹ0 < 1 4&+Š¥+ā&ȗƹɺBȔ '&>ŲǴǌ*,#+ɢç&ǆ)>ɊĆñêČ>ɻ+ƒɜɹéɷɺ*)"%¸7%ǮǸƉ&#+ɢç+ɕ+ïƻǌɝɤBƃµ&>
ɹƟʄFST 'ŒŠ,Ďɞ*,ïƻǌɝɤÿ+Ƈ)jDO[}+ĴɪBÏ%ýÁ>7ɻ+ȠɎ*,Ɵńļțɺ'@&ɻ+ŒŠ,¤Ǿ±+ɊĆɬĪɹ¤Ǿ+&ƴČ+ĔǣɊĆ+È7>¿Öɺ+ŖČ¥+ɖŠ& >7ɻǸȢǌ*Ŵń)ǷżBĹ>7*,ɻ~#+Ŕɢï<āɑ*EJ

Y. Tanabe - 6SBÇɤ%ȠŻ>ļț >&ǥȀ<,+ãɮ*Î=ǵ67ɻ»ƁğƖǮ+ơ
ó*ǊɆ% >șŠ+ƪƛBrawïó*ȤČɻ< 2005 ħþĈ*EJS+āɑŔɢ|¶ɤôɲBȔ ɻ?<¶ɤƀ*# % MLST ȠŻBȔ"ǷżB¬* FST Bƿ ȠŻBȔ"+Ƿżɻ¯ǌ*Ȝ>' FST ,Ŵń*Ǝ+¥BǙ' (FST = 0.351, P < 0.001)ɻ#4=ÕïƮ+¤Ǿɕ+ɊĆǵŇ*ɇ Ȝ<?>'A"ɻ+ɇ +h[}z,Çǰ&,)" (éʂ)Ų8ɴ1Ƿż,ɻA 3 Myɤ?% ) ×~ƪƛ±+ïƮɕ&
EJS+ɊĆǵŇ¯ǆ)"% '& ><*ɴ1'*ɻ×~+ïƮ+EJ
S+ɊĆǵŇ 2 {ųķ*¯ǆ)>8+*ýÂ%  (Tanabe et al. 2009b)

éʂ »ƁğƖǮ+EJS¤Ǿ* >Ɋǌ¶ÂÕï¤Ǿ* >ɊĆñɬĪ¶ĢB MLST ǮǸƉɹÔɺ* >ʂ#+Pw}l¶ɹPw}l A, B, D- Gɺ*õ$ %²Puj&Ǚ²Puj+ɩǟ,¶ɤƀ+¤Š*Ɣ>; *ŗPw}l C ,ėǾ+7ǑǅȗɹÔɺ+ FST ¥+ŴńƖƬ,0.1ɸŲ8ā) FST ¥BǙɹŲ8ǆ)"ɊǌǵŇBǙɺ¤Ǿ+nE,ɻ×~ïƮɹɶĞɻɦ{Ƣɺ+ 7 ų' 9 ųɻË.Ųȼŕ>ɢçɹɶĞ'ɖɻ×ŧ*Ŕɢɺ& >'*Ɵń%ɻåȟē?șɣ)pOyV`FX+¤Ǿ+Ɋǌǐɇh[}zB(+; *ȠɎ?-;@ ʅ MLST +ǮǸƉBǭž%5>'ɻ+#+ńÿ)ȟēǷż*,ɻPw}l G 'Ǜ>Ǟ±ǮǸǾɖ% >'A>ɽǤ&ȫũȵɤ*;>ɝɤ,ɻ»ƁğƖǮ+ƪƛ+pOyVX`FX*%,ȟē?)"ɻPw}l G Bɚ'į )<8ȵɤ*;>ɝɤBǙá>h[}z8Ȝ<? (Tanabe et al. 2009b)ǥȀ<,
+Pw}l G +Ǟ±ǮǸǾɹAʆFɺ',ǆ)>ƽņǌƴŀBő#ɈĽǌ)Pw}l&>ÒȅŀBǿ % =ɻƹî+ƴŀ+ȠŻBɂ7% > ?*%8ɻĘ)'8å+ïƻXQ}wɹ60 MyǝĪɺ*%,ɻïƻǌȵɤ9Ɩó+¶Ť' "ïƻǌțǱÿ+j

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FX,ǃÕï&Ȝ<?>;=ā ïƻXQ}w&ȠŻ÷Ö*ɻ(+; )ǷżĹ<?>,ȋÞƦ '@& >EJS, ”Everyting is everywhere” ȫȽ1>; *ɻǃÕï&; )ɊĆǵŇBő"% >+& @ ʅ?'8ïóêŴ+ǮǸǾȜµ?
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ïó;=ɢǌ*pOyV`FX+Ŕɢ|Çɤ|ɊĆ[GizPBȔ"% >ĕźǌ*,ƍĠɻťāɛɻEjvKǦ*%8ŔɢBȔ ɻ×ǞBŸţ'ïƺȝƈ+ƽƳïƻĉǌǗǠ

BĚɔ 'ǿ % >(!%Avise, J.C. 2000. Phylogeography: The history and formation of species. Cambridge (Massachusetts):
Harvard University Press.
Baas Becking, L.G.M. 1934. Geobiologie of inleiding tot de milieukunde. The Hague, the Netherlands: W.P.
Van Stockum & Zoon (in Dutch).
Hanage, W.P., Fraser, C. & Spratt, B.G. 2005. Fuzzy species among recombinogenic bacteria. BMC Biol. 3:
6.
Janse, I., Kardinaal, W.E., Meima, M., Fastner, J., Visser, P.M. & Zwart, G. 2004. Toxic and nontoxic
Microcystis colonies in natural populations can be differentiated on the basis of rRNA gene internaltranscribed spacer diversity. Appl. Environ. Microbiol. 70: 3979-3987.
Maiden, M.C., Bygraves, J.A., Feil, E., Morelli, G., Russell, J.E., Urwin, R., Zhang, Q., Zhou, J., Zurth, K.,
Caugant, D.A., Feavers, I.M., Achtman, M. & Spratt B.G.. 1998. Multilocus sequence typing: a portableapproach to the identification of clones within populations of pathogenic microorganisms. Proc. Natl.
Acad. Sci. USA 95: 3140-3145.
Otsuka, S., Suda, S., Shibata, S., Oyaizu, H., Matsumoto, S. & Watanabe M.M. 2001. A proposal for theunification of five species of the cyanobacterial genus Microcystis Kutzing ex Lemmermann 1907 under
the rules of the Bacteriological Code. Int. J Syst. Evol. Microbiol. 51: 873-879.
Papke, R.T., Ramsing, N.B., Bateson, M.M. & Ward D.M. 2003. Geographical isolation in hot spring
cyanobacteria. Environ. Microbiol. 5: 650-659.
Papke, R.T. & Ward, D.M. 2004. The importance of physical isolation to microbial diversification. FEMS
Microbiol. Ecol. 48: 293-303.
Slatkin, M. 1993. Isolation by distance in equilibrium and non-equilibrium populations. Evolution 47:
264-279.
Smith, J.M., Smith, N.H., O'Rourke, M. & Spratt, B.G. 1993. How clonal are bacteria? Proc. Natl. Acad. Sci.
USA 90: 4384-4388.
Tanabe, Y., Kaya, K. & Watanabe, M.M. 2004. Evidence for recombination in the microcystin synthetase(mcy) genes of toxic cyanobacteria Microcystis spp. J. Mol. Evol. 58: 633-641.
Tanabe, Y., Kasai F. & Watanabe, M.M. 2007. Multilocus sequence typing (MLST) reveals high genetic
diversity and clonal population structure of the toxic cyanobacterium Microcystis aeruginosa.

Y. Tanabe - 8

Microbiology 153: 3695‫‏‬-3703.
Tanabe, Y., Sano, T., Kasai, F. & Watanabe, M.M. 2009a. Recombination, cryptic clades and neutral
molecular divergence of the microcystin synthetase (mcy) genes of toxic cyanobacterium Microcystis
aeruginosa. BMC Evol. Biol. 9: 115.
Tanabe, Y., Kasai, F. & Watanabe, M.M. 2009b. Fine-scale spatial and temporal genetic differentiation ofwater bloom-forming cyanobacterium Microcystis aeruginosa: revealed by multilocus sequence typing.
Environ. Microbiol. Reports 1: 575-582.
Tillett, D., Parker, D.L. & Neilan, B.A. 2001. Detection of toxigenicity by a probe for the microcystin
synthetase A gene (mcyA) of the cyanobacterial genus Microcystis: comparison of toxicities with 16S
rRNA and phycocyanin operon (phycocyanin intergenic spacer) phylogenies. Appl. Environ.
Microbiol .67: 2810-2818.
Whitaker, R.J., Grogan, D.W. & Taylor, J.W. 2003. Geographic barriers isolate endemic populations ofhyperthermophilic archaea. Science 301: 976-978.
Whitaker, R.J. 2006. Allopatric origins of microbial species. Philos. Trans. R. Soc. Lond. B Biol. Sci. 361:
1975-1984.

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